1 The global distribution of tetrapods reveals a need for targeted reptile 2 conservation 3 4 Uri Roll#1,2, Anat Feldman#3, Maria Novosolov#3, Allen Allison4, Aaron M. Bauer5, Rodolphe 5 Bernard6, Monika Böhm7, Fernando Castro-Herrera8, Laurent Chirio9, Ben Collen10, Guarino R. 6 Colli11, Lital Dabool12 Indraneil Das13, Tiffany M. Doan14, Lee L. Grismer15, Marinus 7 Hoogmoed16, Yuval Itescu3, Fred Kraus17, Matthew LeBreton18, Amir Lewin3, Marcio Martins19, 8 Erez Maza3, Danny Meirte20, Zoltán T. Nagy21, Cristiano de C. Nogueira19, Olivier S.G. 9 Pauwels22, Daniel Pincheira-Donoso23, Gary Powney24, Roberto Sindaco25, Oliver Tallowin3, 10 Omar Torres-Carvajal26, Jean-François Trape27, Enav Vidan3, Peter Uetz28, Philipp Wagner5,29, 11 Yuezhao Wang30, C David L Orme6, Richard Grenyer✝1 and Shai Meiri✝*3 12 13 # Contributed equally to the paper 14 ✝ Contributed equally to the paper 15 * Corresponding author 16 17 Affiliations: 18 1 School of Geography and the Environment, University of Oxford, Oxford, OX13QY, UK. 19 2 Mitrani Department of Desert Ecology, The Jacob Blaustein Institutes for Desert Research, 20 Ben-Gurion University, Midreshet Ben-Gurion 8499000, Israel. (Current address) 21 3 Department of Zoology, Tel-Aviv University, Tel-Aviv 6997801, Israel. 22 4 Hawaii Biological Survey, 4 Bishop Museum, Honolulu, HI 96817, USA. 23 5 Department of Biology, Villanova University, Villanova, PA 19085, USA. 24 6 Department of Life Sciences, Imperial College London, Silwood Park Campus Silwood Park, 25 Ascot, Berkshire, SL5 7PY, UK 26 7 Institute of Zoology, Zoological Society of London, London NW1 4RY, UK. 27 8 School of Basic Sciences, Physiology Sciences Department, Universidad del Valle, Colombia. 28 9 14, rue des roses - 06130 Grasse, France. 29 10 Centre for Biodiversity & Environment Research, University College London, London WC1E 30 6BT, UK. 31 11 Departamento de Zoologia, Universidade de Brasília, 70910-900, Brasília, Distrito Federal, 32 Brazil. 33 12 Department of Genetics and Developmental Biology, The Rappaport Family Institute for 34 Research in the Medical Sciences, Faculty of Medicine, Technion – Israel Institute of 35 Technology, Haifa 31096, Israel. 36 13 Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 37 94300 Kota Samarahan, Sarawak, Malaysia. 38 14 Department of Biology, University of Central Florida, Orlando, FL 32816, USA. 39 15 Department of Biology, La Sierra University, Riverside, CA 92505, USA. 40 16 Museu Paraense Emílio Goeldi/CZO, Caixa Postal 399, 66017–970 Belém, Pará, Brazil. 41 17 Department of Ecology and Evolutionary Biology, University of Michigan, Ann-Arbor, MI 42 48109-1048, USA. 43 18 Mosaic, (Environment, Health, Data, Technology), Yaoundé, Cameroon. 44 19 Departamento de Ecologia, Instituto de Biociências, Universidade de São Paulo, 05508-090 45 São Paulo, São Paulo, Brasil. 46 20 Royal Museum for Central Africa, Leuvensesteenweg 13, 3080 Tervuren, Belgium. 47 21 Joint Experimental Molecular Unit, Royal Belgian Institute of Natural Sciences, B-1000 48 Brussels, Belgium. 49 22 Département des Vertébrés Récents, Royal Belgian Institute of Natural Sciences, B-1000 50 Brussels, Belgium. 51 23 School of Life Sciences, Joseph Banks Laboratories, University of Lincoln, Brayford Campus, 52 Lincoln, LN6 7DL, UK. 53 24 NERC Centre for Ecology and Hydrology, Maclean Building, Crowmarsh Gifford, 54 Wallingford, OX10 8BB, UK. 55 25 Museo Civico di Storia Naturale, I-10022 Carmagnola (TO), Italy. 56 26 Museo de Zoología, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del 57 Ecuador, Apartado 17-01-2184, Quito, Ecuador. 58 27 Institut de Recherche pour le Développement, Laboratoire de Paludologie et Zoologie 59 Médicale, UMR MIVEGEC, Dakar, Senegal. 60 28 Center for the Study of Biological Complexity, Virginia Commonwealth University, 61 Richmond, VA 23284, USA. 62 29 Zoologische Staatssammlung München, D-81247 München, Germany. 63 30 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China. 64 Abstract 65 The distributions of amphibians, birds and mammals have underpinned global and local 66 conservation priorities, and have been fundamental to our understanding of the determinants of 67 global biodiversity. In contrast, the global distributions of reptiles, representing a third of 68 terrestrial vertebrate diversity, have been unavailable. This prevented reptiles’ incorporation into 69 conservation planning and biased our understanding of the underlying processes governing 70 global vertebrate biodiversity. Here, we present and analyse, for the first time, the global 71 distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness 72 patterns of the other three tetrapod classes are good spatial surrogates for species richness of all 73 reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. 74 Hotspots of total and endemic lizard richness overlap very little with those of other taxa. 75 Moreover, existing protected areas, sites of biodiversity significance and global conservation 76 schemes, represent birds and mammals better than reptiles. We show that additional conservation 77 actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile 78 knowledge to a global complementarity conservation priority scheme, identifies many locations 79 that consequently become important. Notably, investing resources in some of the world’s arid, 80 grassland, and savannah habitats might be necessary to represent all terrestrial vertebrates 81 efficiently. 82 Introduction 83 Our knowledge of the distributions of a broad variety of organisms has improved greatly in the 84 past decade1-3. This has greatly aided our efforts to conserve biodiversity4-6 and significantly 85 enhanced our grasp of broad scale evolutionary and ecological processes7-12. Nevertheless, 86 despite comprising one third of terrestrial vertebrate species, knowledge of reptile distributions 87 remained poor and unsystematic. This represented a major gap in our understanding of the global 88 structure of biodiversity and our ability to conserve nature. Historically, broad-scale efforts 89 towards the protection of land vertebrates (and thus also of reptiles) have been based 90 predominantly on data from plants, birds, mammals and to a lesser degree amphibians13-15. Here 91 we present complete species-level global distributions of nearly all reptiles: 10,064 known, 92 extant, terrestrial species for which we could identify precise distribution information. These 93 distributions cover the Sauria (lizards, 6110 species), Serpentes (snakes, 3414 species), 94 Testudines (turtles, 322 species), Amphisbaenia (‘worm lizards’, 193 species), Crocodylia 95 (crocodiles, 24 species) and Rhynchocephalia (the tuatara, one species). 96 This dataset completes the global distribution mapping of all described, extant, terrestrial 97 vertebrates (Fig. 1a), providing information that has been missing from much of the global 98 conservation planning and prioritization schemes constructed over the last twenty years4. We use 99 our reptile distribution data to: a) examine the congruence in general, hotspot, and endemism 100 richness patterns across all tetrapod classes and among reptile groups; b) explore how current 101 conservation networks and priorities represent reptiles; and c) suggest regions in need of 102 additional conservation attention to target full terrestrial vertebrate representation and highlight 103 current surrogacy gaps, using a formal conservation prioritisation technique. 104 105 Results and Discussion 106 Species richness of reptiles compared to other tetrapods 107 The global pattern of reptile species richness (Fig. 1b) is largely congruent with that of all other 108 terrestrial vertebrates combined (r = 0.824, e.d.f. = 31.2, p << 0.0001; Figs. 2a, S1, Table S1). 109 However, the major reptile groups (Figs. 1c-e, 2b-c, S1, Table S1) show differing degrees of 110 congruence with the other tetrapod taxa. The richness distribution of snakes (Fig. 1d) is very 111 similar to that of other tetrapods (Fig. 2c) in showing pan-tropical dominance (r = 0.873, e.d.f. = 112 30.2, p << 0.0001). Lizard richness is much less similar to non-reptilian tetrapod richness (r = 113 0.501, e.d.f. = 38.3, p << 0.001, Fig. 2b). It is high in both tropical and arid regions, and notably 114 in Australia (Figs. 1c, S1). Turtle richness is also less congruent with diversity patterns of the 115 other tetrapods (r = 0.673, e.d.f. = 55.2, p << 0.001), and peaks in the south-eastern USA, the 116 Ganges Delta, and Southeast Asia (Fig. 1e). 117 Snakes dominate reptile richness patterns due to their much larger range sizes compared to 118 lizards, even though lizards are about twice as speciose (median ranges size for 3414 snake 119 species: 62,646 km2; for 6415 lizard species: 11,502 km2; Fig. S2). Therefore snakes, 120 disproportionally influence global reptile richness patterns16,17 (Table S1, Fig. S1). 121 Hotspots of richness and range-restricted species 122 As with overall richness patterns, hotspots of richness (the richest 2.5%, 5%, 7.5% and 10% of 123 grid-cells) for all reptiles combined, and of snakes, are largely congruent with those of other 124 tetrapod classes. However they are incongruent with hotspots of lizard or turtle richness (Figs. 3; 125 S3). 126 Congruence in the richness of range-restricted species (those species with the smallest 25% or 127 10% ranges in each